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Rapid Evolution of the Visual System: a Cellular Assay of the Retina and Dorsal Lateral Geniculate Nucleus of the Spanish Wildcat and the Domestic Cat

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Rapid Evolution of the Visual System: a Cellular Assay of the Retina and Dorsal Lateral Geniculate Nucleus of the Spanish Wildcat and the Domestic Cat The Journal of Neuroscience, January 1993, 13(l): 208-229 Rapid Evolution of the Visual System: A Cellular Assay of the Retina and Dorsal Lateral Geniculate Nucleus of the Spanish Wildcat and the Domestic Cat Robert W. Williams,’ Carmen Cavada,2 and Fernando Reinoso-Suhrez* ‘Department of Anatomy and Neurobiology, College of Medicine, University of Tennessee, Memphis, Tennessee 38163 and *Departamento de Morfologia, Facultad de Medicina, Universidad Aut6noma de Madrid, 28029 Madrid, Spain The large Spanish wildcat, Fe/is silvestris tartessia, has re- and important topic, it has been difficult to study the process tained features of the Pleistocene ancestor of the modern of brain evolution in any detail. Our approach has been to domestic cat, F. catus. To gauge the direction and magnitude identify a pair of closely related living species,one from a highly of short-term evolutionary change in this lineage, we have conservative branch that has retained near identity with the compared the retina, the optic nerve, and the dorsal lateral ancestral species,and the other from a derived branch that has geniculate nucleus (LGN) of Spanish wildcats and their do- undergone rapid evolutionary change. The recent recognition mestic relatives. Retinas of the two species have the same that evolution and speciationcan occur in short bursts separated area. However, densities of cone photoreceptors are higher by long interludes of stasisprovides a sound theoretical basis in wildcat-over 100% higher in the area centralis-where- for a search for such pairs (Schindewolf, 1950; Eldredge and as rod densities are as high, or higher, in the domestic lin- Gould, 1972; Stanley, 1979; Gould and Eldredge, 1986). eage. Densities of retinal ganglion cells are typically 20- A candidate pair among mammals is the wildcat, Fe/is sil- 100% higher across the wildcat retina, and the total ganglion vestris, and the domestic cat, F. catus. Historical, biochemical, cell population is nearly 70% higher than in the domestic and genetic data point to a derivation of the domestic cat from cat. These differences are confined to the populations of the North African wildcat, F. silvestris lybica (Darwin, 1890; beta and gamma retinal ganglion cells. In contrast, the pop- Clutton-Brock, 1981; Benveniste, 1985; Collier and O’Brien, ulation of alpha cells is almost precisely the same in both 1985; Wayne et al., 1989). This small North African wildcat species. The wildcat LGN is much larger than that of the descendedfrom a larger ancestorwhose Pleistocene fossil record domestic cat and contains-50% more neurons. However, has been described carefully by Km-ten (1965a,b). These Pleis- cell size does not differ appreciably in either the retina or tocene wildcats share a constellation of features-short pre- LGN of these species. The differences in total numbers of molar-canine diastema, large canine and carnassialteeth, and ganglion cells and LGN neurons correspond neatly to the a low crowned fourth lower premolar-with the particular sub- overall decline in brain size in the domestic lineage and to specieswe have studied, F. silvestristartessia (Km-ten, 1965a,b). allometric predictions based on average species differences In fact, the Spanishwildcat has more in common with wildcats in body size. We suggest that an increase in the severity of living toward the end of the Wtirm glaciation, 15,000-20,000 naturally occurring cell death is the most plausible mecha- years before present (BP) than with other extant F. silvestris nism that can account for the rapid evolutionary reduction subspecies(Table 1). For this reason,KurtCn (1965b) concluded in cell populations in this feline lineage. that the Spanish subspeciesis an isolated relict of an ancient [Key words: brain size, allometry, rods and cones, retinal Mediterranean wildcat population that has survived with min- ganglion cell, optic nerve, lateral geniculate nucleus, cell imal change for over 20,000 years in a warm Iberian refuge. death] We decided that a comparison of this particular wildcat sub- specieswith the domestic cat could provide detailed insight into The tempo of mammalian brain evolution has beenrapid. Two- the range, rate, and direction of evolutionary change in brain fold changesin brain weight have occurred over periods of l- structure. For example, we thought we would be able to deter- 5 million years in several lineages,including that of humans mine whether the rapid decline in body size in the cat lineage (Edinger, 1948, 1966; Jerison, 1973, 1979; Radinsky, 1973, has been associatedwith a matched decline in brain massand, 1975, 198 1; Eisenberg, 1981). Although this is an interesting if so, whether this changehas been brought about by a reduction in neuron number or an increase in cell packing density. We Received Apr. 14, 1992; revised July 13, 1992; accepted July 15, 1992. also thought it would be possibleto determine whether numer- This research was supported in part by Grant DGICYT PB88-0170 and a grant ical relations among interconnected parts of the brain have been from the NIH. We are especially indebted to Mariano Sanz Pech and Antonio conserved in this rapidly evolving mammalian lineage, and if Ptrez Alonso-Geta ofthe Institute for the Conservation ofNature, Madrid, Spain, so, then how precisely. Finally, we thought it might be possible for wildcat material. We thank Drs. Leo Chalupa, Andrea Elberger, and Douglas Frost for the generous use of material from their collections of domestic cat tissue, to compare ontogenetic differencesin thesespecies and thereby and Drs. Leo Chalupa, Robert Foehring, Dan Goldowitz, Todd Preuss, Anton expose developmental and cellular mechanismsthat ultimately Reiner, and Michael Rowe for critical comments on drafts of the manuscript. Correspondence should be addressed to Dr. Robert Williams at the above underlie phyletic change. address. Taking advantage of the wealth of data on the structure of Copyright 0 1993 Society for Neuroscience 0270-6474/93/130208-21$05.00/O the primary visual systemof domestic cats, we have begunwith The Journal of Neuroscience, January 1993, 13(i) 209 a quantitative analysis of populations of cells in the retina and Femurs of European wildcats are typically less than 13 cm long (Rijhrs, the dorsal lateral geniculate nucleus (LGN) of the Spanish wild- 1955). HRP injection procedure. We made injections of horseradish perox- cat. In this analysis we make two explicit assumptions: first, idase (HRP) into the left dorsal lateral geniculate nucleus (LGN), left that th; adult wildcats, which we have had the very rare op- superior colliculus, and left pretectal region of the male wildcat (SI). portunitv to studv,_ I are representative of the Spanish subspecies; Similar unilateral (n = 1) and bilateral (n = 3) series of injections were and second, that the Spanish wildcat is itselfrepresentative of, made into normal domestic cats. Some of the domestic cat retinas have or at least similar to, the ancestral population of Pleistocene been described and illustrated in previous studies (Chalupa et al., 1984; Kirby and Chalupa, 1986). Animals initially received a single intra- wildcats from which domestic cats ultimately trace their descent. muscular dose of ketamine f0.4~ cc)I and atrooine (0.2 cc) and were The validity of these key assumptions is supported by data subseauentlv anesthetized with an intravenous miection of sodium pen- summarized in the next section (Table 1). From this vantage tobarbital (30 mg/kg). They were placed in a stereotaxic device-and point, we attempt to assess structural changes in the primary craniotomies were made over the approximate locations of the left superior colliculus, pretectum, and dorsal lateral geniculate nucleus. The visual system associated with approximately 25,000 years of boundaries of retinorecipient nuclei were mapped by recording multi- natural selection at the close of the Pleistocene, followed by cellular visually evoked activity with a tungsten microelectrode..A series 3000-4000 years of domestication. of 150-200 nl iniections of 25% HRP diluted in oure dimethvl sulfoxide (DMSO) were made along each penetration within the LGN (23 pen- etrations), pretectum (6 penetrations), and superior colliculus (10 pen- etrations) as described in Chalupa et al. (1984). The total volume of Materials and Methods HRP delivered in a single penetration ranged from 200 to 600 nl. We received two adult wildcats (Felis silvestris tartessiu Miller), a male Perfusion and dissection of the animals. All animals received an over- (SI) and a female (S2), from the Institute for the Conservation ofNature dose of pentobarbital(200 mg) and were subsequently perfused through (ICONA), Spain. Both animals had been rescued from game traps in the heart with phosphate-buffered saline (0.1 M, pH 7.3), followed by a the Sierra Morena of south central Spain. The two thin but healthy mixture of 1.25% paraformaldehyde and 2.5% glutaraldehyde in phos- animals had been held for several months in a large outdoor pen. Breed- phate buffer. We removed the cranial vault in one large intact piece, ing was thought to have failed. Both animals had the coat color and and the brain-including olfactory bulb, pituitary, and the entire me- pattern typical of wildcats and many domestic tabby cats (Haltenorth, dulla (Fig. I)-the eyes, and segments of most cranial nerves were re- 1953; Rodriguez de la Fuente, 1979). Each had four prominent dark moved. We took retinas from the eyes and flattened them by making a streaks running down the neck, a single broad dark band running along set of radial cuts. The areas of flat-mounted retinas were measured before the dorsal midline to the base of the tail, six to eight less distinct dark dehydration. We cleaned crania and dentaries in a 25% solution of arched bands running from back to abdomen, and three to four dark bleach. Pelts of both wildcats were returned to ICONA. rings around limbs and tails. Figure 26 in Haltenorth (1953) shows a Following the transcardial perfusion of wildcat S2, an adult female specimen of the F.
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